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Historical BiologyAn International Journal of Paleobiology

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A new sauropodomorph ichnogenus from theLower Jurassic of Sichuan, China fills a gap in thetrack record

Lida Xing, Martin G. Lockley, Jianping Zhang, Hendrik Klein, Daqing Li,Tetsuto Miyashita, Zhongdong Li & Susanna B. Kümmell

To cite this article: Lida Xing, Martin G. Lockley, Jianping Zhang, Hendrik Klein, Daqing Li,Tetsuto Miyashita, Zhongdong Li & Susanna B. Kümmell (2016) A new sauropodomorphichnogenus from the Lower Jurassic of Sichuan, China fills a gap in the track record, HistoricalBiology, 28:7, 881-895, DOI: 10.1080/08912963.2015.1052427

To link to this article: http://dx.doi.org/10.1080/08912963.2015.1052427

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A new sauropodomorph ichnogenus from the Lower Jurassic of Sichuan, China fills a gap in thetrack record

Lida Xinga*, Martin G. Lockleyb, Jianping Zhanga, Hendrik Kleinc, Daqing Lid, Tetsuto Miyashitae, Zhongdong Lif and

Susanna B. Kümmellg

aSchool of the Earth Sciences and Resources, China University of Geosciences, Beijing 100083, China; bDinosaur Trackers ResearchGroup, University of Colorado at Denver, CB 172, PO Box 173364, Denver, CO 80217-3364, USA; cSaurierwelt PaläontologischesMuseum, Alte Richt 7, D-92318, Neumarkt, Germany; dGeological Museum of Gansu, Lanzhou 730040, China; eDepartment ofBiological Sciences, University of Alberta, 11455 Saskatchewan Drive, Edmonton, Alberta T6G 2E9, Canada; fGeophysical Team ofSichuan Bureau of Geological and Mineral Investigation and Exploration, Chengdu, Sichuan 610072, China; gInstitute of EvolutionaryBiology, University Witten/Herdecke, Stockumerstr. 10-12, 58454 Witten, Germany

(Received 8 January 2015; accepted 14 May 2015)

A dinosaur tracksite in the Lower Jurassic Ziliujing Formation of Sichuan Province, China consists of a spectacular sub-vertical exposure, with multiple track-bearing levels and trackways showing parallel and bimodal orientations. Based onwell-preserved material, the new ichnogenus and ichnospecies, Liujianpus shunan ichnogen. nov. ichnosp. nov. is erected toaccommodate distinctive sauropodomorph trackways occurring in this assemblage. Liujianpus has a unique combination offeatures, some relating to the early Jurassic basal sauropodomorph (prosauropod in traditional usage) ichnogenus Otozoum,others to the sauropod ichnogenus Brontopodus. Despite such a mix of basal sauropodomorph- and sauropod-like features,the trackmaker of Liujianpus is likely a basal sauropodomorph. This identification is consistent with the occurrence of basalsauropodomorph skeletons from geographically and chronologically close localities. The other distinct morphotype from thetracksite is linked to a sauropod trackmaker. As such, the ichnofauna consisting of two distinct foot morphotypes reflects thediversity of sauropodomorph dinosaurs in the Early Jurassic of Asia.

Keywords: Sauropodomorph tracks; Liujianpus; Ziliujing Formation; Lower Jurassic; Sichuan Province; China

1. Introduction

Although skeletal remains of sauropodomorph dinosaurs

are well known, and locally abundant in the early

Mesozoic of East Asia (Young 1951; Dong 1992; Barrett

et al. 2005), their tracks are scarcely known (Lockley et al.

2002) and have not been described in detail. Documented

tracks were mostly attributed to sauropods, whereas

diagnostic basal sauropodomorph tracks have not been

reported from East Asia thus far. This situation is in

contrast to the track record of other regions, notably North

America, Europe and southern Africa, where the basal

sauropodomorph track record is well documented and of

considerable significance in the history of ichnology.

1.1. Global distribution of basal sauropodomorphtracks

The first ichnogenus currently confidently attributed to

prosauropods was Otozoum, which was named by

Hitchcock (1847) from the Lower Jurassic Portland

Sandstone of New England. However, Hitchcock did not

link Otozoum with a prosauropod trackmaker since that

group was unknown at the time. Subsequently, Lull (1904,

1953) assigned Otozoum as a prosauropod track and

erected the ichnofamily Otozoidae in a revision of

Hitchcock (1847, 1848, 1858). Recent work by Rainforth

(2003), Lockley et al. (2006a) and others unanimously

support the basal sauropodomorph affinity for Otozoum.

Ellenberger (1970, 1972, 1974) named the ichnogenus

Pseudotetrasauropus from the Lower Jurassic of southern

Africa. Features similar to Otozoum let some workers infer

a basal sauropodomorph affinity for this ichnotaxon

(Ellenberger and Ellenberger 1958; Ellenberger et al.

1969; Ellenberger 1970, 1972; Lockley and Meyer 2000;

D’Orazi Porchetti and Nicosia 2004, 2007 ; Lockley et al.

2006a). However, Rainforth (2003) considered a crur-

otarsan origin based on the derived phalangeal formula.

Olsen and Galton (1984) interpreted Pseudotetrasauropus

as a bipedal Brachychirotherium suggesting a crocodylian-

stem affinity as well. The latter authors revised the

ichnotaxonomy of Pseudotetrasauropus and other

southern African ichnogenera based on a survey of the

literature, but without first-hand observations of the

original African materials. Although they synonymised

many of the southern African ichnogenera with North

American ichnotaxa, Otozoum and Pseudotetrasauropus

are now considered distinct (Lockley and Meyer 2000;

Rainforth 2003; D’Orazi Porchetti and Nicosia 2004;

Lockley et al. 2006a; Gierliński, 2009).

q 2015 Taylor & Francis

*Corresponding author. Email: xinglida@gmail.com

Historical Biology, 2016

Vol. 28, No. 7, 881–895, http://dx.doi.org/10.1080/08912963.2015.1052427

mailto:xinglida@gmail.commailto:xinglida@gmail.comhttp://dx.doi.org/10.1080/08912963.2015.1052427

At present, the plexus of tracks attributed to basal

sauropodomorphs includes Otozoum, Pseudotetrasauropus,

Evazoum and Kalosauropus and has been dubbed the

OPEK plexus (Lockley et al. 2006a), which belongs to the

ichnofamily Otozoidae (Lockley et al. 2006a; Lockley and

Lucas 2013). Lavinipes cheminii described by Avanzini

et al. (2003) could probably also be assigned to this plexus

since it is generally similar to Otozoum, although not

preserved with sufficient detail to show the complex pads so

characteristic of Otozoum. The former two OPEK plexus

ichnogenera (Otozoum and Pseudotetrasauropus) are based

on large type specimens, and include large robust forms that

represent quadrupedal as well as bipedal trackmakers.

In contrast, the two latter ichnogenera represent small

bipeds, which were probably gracile animals.

In contrast to the long history of research on basal

sauropodomorph tracks, the study of true sauropod tracks

did not begin until the 1930s (Bird 1939, 1985) and it was

not until 1989 that the ichnogenus Brontopodus was

named for large sauropod tracks from the Cretaceous of

Texas (Farlow et al. 1989). Since then, most additional

sauropod track reports, as well as newly proposed

ichnogenera have pertained to late Mesozoic (Late

Jurassic-Late Cretaceous) tracks.

Although sauropods are known from the Late Triassic

(Buffetaut et al. 2000, 2002), there have been relatively few

reports of sauropod or sauropodomorph tracks from Triassic

and Lower/Middle Jurassic strata, and few of these are

represented by newly named ichnotaxa. The oldest distinctive

sauropodomorph ichnogenus attributable to a robust quad-

ruped is Eosauropus (Lockley et al. 2006b) named for well-

preserved material from the Late Triassic of New Mexico

(previously informally referred to the ichnogenus Tetrasaur-

opus; Ellenberger 1972). In North America, Eosauropus is

often represented by poorly preserved trackways, presumably

made under substrate conditions not suitable for optimal

preservation (Lockley et al. 2011). One set of tracks

resembling poorly preserved Eosauropus (or Otozoum) has

been tentatively identified from China (Xing et al. 2014a).

Other reports of Early and Middle Jurassic sauropod

tracks come fromMorocco (Ishigaki 1988; Gierliński et al.

2009), Italy (e.g. Leonardi and Mietto 2000), Poland (e.g.

Gierliński et al. 2004; Gierliński, 2009) and Portugal

(Santos et al. 1994), respectively. In the former case, the

tracks, which remain unnamed, are Early Jurassic in age

and resemble those described here, whereas in the later

case, the tracks (Polyonyx gomesi) are Middle Jurassic in

age and are morphologically distinct from any Lower

Jurassic forms (Santos et al. 2009). Only two previous

reports of sauropod tracks from the Lower Jurassic of

China are known: (a) from the Zhenzhucheng Formation

of the Dazu region of Sichuan (Matsukawa et al. 2006)

illustrated by Lockley and Matsukawa (2009); and (b)

from the Ziliujing Formation of Zigong City, Sichuan

(Xing et al. 2014b).

1.2. Sauropodomorph trackways from Gulin County

At the southern border of the Sichuan Basin, the Gulin area

protrudes into northern Guizhou Province as a peninsular-

shaped region. In May 2009, Yiguang Chen and Jianming

Tang, the engineers from No. 113 Geological Team,

Sichuan Bureau of Geology and Mineral Resources,

discovered a group of dinosaur tracks at Heping (meaning

‘peace’) brickfield near Zhongshan Village, Jiaoyuan

(meaning ‘pepper garden’) Township, Gulin County.

During 2010–2014, the senior author has made six visits

to investigate the tracksite. The purpose of this paper is to

describe a large assemblage of sauropodomorph trackways

from a locality near Gulin County in the Lower Jurassic

Ziliujing Formation in Sichuan Province, China, and

discuss their ichnotaxonomic status and paleobiological

significance. This paper elaborates on a very preliminary

note on the site by Xing (2010). Co-occurring theropod

tracks and trackways from the same locality are described

in a different study (Xing, Lockley, Zhang et al. in press).

Institutional and other abbreviations: JY ¼Jiaoyuan tracksite; UCM ¼ University of ColoradoMuseum of Natural History, Boulder, Colorado, USA;

ZLJ ¼ World Dinosaur Valley Park, Yunnan Province,China; S ¼ Sauropod; I ¼ Isolated; M ¼ Manus; P ¼ Pes;L ¼ Left; R ¼ Right

2. Geographic and geologic setting

The Jiaoyuan tracksite is situated at latitude N 24802032.7800 and longitude E 105848053.0500 in a small, relativelyremote valley 30 km south southeast of Gulin County, in

Sichuan Province (Figure 1). The main tracksite consists

of an outcrop consisting of bedding planes that dip steeply

at 708 to the southeast (Figure 2). Quarrying operations toextract siltstones and mudstones for a local brick factory

have ceased now. The main track-bearing surface

approximately faces south. Long parallel trackways are

conspicuous, especially in the western sector of the

exposed surface where the upper exposures are to the north

and the lower exposures to the south. Close inspection of

this main western outcrop reveals that some of these

trackways extend in opposite directions indicating a

bimodal orientation to the overall assemblage (Figure 3).

This western sector is divided from an eastern sector by

local outcrops of the overlying beds. As noted further in

the text, a few accessible tracks in the lower part of the

eastern sector have been studied. At the base of the

western outcrop, this is a relatively small exposure of a

track-bearing sandstone unit overlying the main surface.

A single sauropodomorph trackway is exposed on the

surface of this overlying unit.

Heim (1930) referred the lower part of the red beds of

the Sichuan Basin to the ‘Ziliujing Series’. When

classifying the strata of eastern Sichuan, Yi (1958)

2 L. Xing et al.882

referred the beds as the ‘Ziliujing Group’. In 1979, the

term Ziliujing Formation was proposed for the lower

Zhenzhuchong Member. Meanwhile, the upper

Lianggaoshan Member was designated as the Dongyue-

miao, Maanshan and Daanzhai Members (Dong 1980).

The Jiaoyuan tracksite is in the Lower Jurassic

Daanzhai Member of the Ziliujing (former spelling:

Tseliutsing) Formation (Gu et al. 1997). The Ziliujing

Formation rests on the Lower Jurassic Zhenzhuchong

Formation and is conformably overlain by the Middle

Jurassic Xintiangou Formation.

The Ziliujing Formation is characterised by the

occurrence of abundant small-sized bivalves dominated

by Apseudocardinia (Cai and Liu 1978). Based on the

presence of Unio huangbogouensis, U. lufengensis,

U. ningxiaensis and Cuneopsis jingyouensis (Ma 1984),

the formation is inferred to be Lower Jurassic in age. The

sporopollenin and ostracod fossils also support this

assessment (Peng et al. 2005).

The Lufengosaurus vertebrate fauna found in the

Daanzhai Member of the Ziliujing Formation includes

Lufengosaurus sp. from the Zigong area, Sichuan Province

(Dong 1984) and the Weixin area, Yunnan Province (The

first division of The Regional Geological Survey Team,

Guizhou Geological Bureau 1979). The Middle Jurassic

Xintiangou Formation contains fragmentary sauropod

material, but basal sauropodomorph body fossils have not

been found in the formation at the time of writing (Peng

et al. 2005). However, although basal sauropodomorph

body fossils mostly flourished in the Lower Jurassic strata

of Asia, only Yunnanosaurus youngi occurs in the Middle

Jurassic (Lu et al. 2007).

The main tracksite occurs as part of a sequence

dominated by friable siltstones and resistant sandstones

(Figure 4), in which a minimum of eight track-bearing

levels have been identified. The measured section has a

total thickness of almost 100 metres, in which there are

only two relatively well-developed sandstone units:

a lower one at the base of the exposure, about 4 metres

thick, and not well exposed, in which tracks could not be

identified, and a well-exposed, upper unit, also about

4 metres thick. These are referred to informally as the

lower and upper sandstone units, with the upper units

Figure 2. Photograph of the Jiaoyuan tracksite showing aspectacular deeply dipping surface with conspicuous parallel andbimodally distributed trackways. A and B corresponding to partof the JYS1 and JYS3 trackways (Figure 5), and JYS11 trackway(Figure 8), respectively.

Figure 1. Map showing the location of the Jiaoyuan tracksite near Gulin in Sichuan Province China.

Historical Biology 3883

being of particular importance because of the rich track

assemblage. All other sandstone units are between 5 and

50 cm in thickness, resulting in a total thickness of

sandstones of about 10 metres, or ,10% of the totalsection. Some surfaces reveal ripple marks and desiccation

cracks.

3. Methods

Due to the steepnessof the main tracksite (,708 dip to thesouth), two experienced climbers were employed to set

ropes and assist with gaining access to the site. This

permitted examination of the tracks at close quarters and

enabled chalk outlines, tracings, photographs and molds to

be made. Once outlines of the tracks in the uppermost

sector of the outcrop had been chalked, a large sheet of

transparent plastic was used to trace most of the trackway

segments in this sector. Due to the size of the outcrop, it

was not feasible to map (trace) the configuration of

trackways across the entire outcrop, especially in the lower

sector where the trackways are longer, therefore portions

of the map were filled in using photographs taken by Xing

(2010).

During our field expedition in August 2013, four

trackway segments (JYSI–IV) in the upper part of the

tracksite were mapped, and the procedure was repeated

again in the fall of 2013. In addition, a further method was

employed on the poorly preserved lower trackways. Using

a professional Canon SLR camera EOS-1D X and super-

telephoto lens EF 800mm f/5.6L IS USM the site was

photographed perpendicular to the surface from the

terraced fields opposite the tracksite. The photographs

were merged using Adobe Photoshop CS6, and the

distribution pattern was mapped.

Ideally, this type of site should be accurately mapped

using Lidar Scanning, photogrammetry or other similar

methods capable of producing three-dimensional images of

large areas. However, obtaining such images is only possible

with specialised equipment and access to a suitable location

or platform from which to operate the scanning and

photogrammetic equipment. Under ideal circumstances,

photogrammetric images might be obtained if a vantage

point such as a balloon, platform or helicopter were to be

situated perpendicular to the surface a few tens of metres

away. Such positioning was practically impossible at such a

remote site, where clear weather is atypical. Moreover,

photogrammetic imaging is compromised by shadows in the

tracks. Even if the above was possible, in order to adequately

describe the tracks, it is still necessary to climb on the

surface to set scales, reference points, inspect details of track

morphology, obtain close-up photographs and moulds.

Using tracings obtained from both the main surface,

and a second surface one metre above, we obtained

following measurements of manus and pes imprints and

trackways: Track length and width, rotation, pes–pes

and manus–manus pace angulation, step, stride and inner

and outer trackway width. The sauropodomorph trackway

parameters follow those of Marty (2008, Figures 2.10 and

2.11). Trackways were numbered JYS1 to JYS10 with

‘JY’ indicating the Jiaoyuan tracksite and ‘S’ the sauropod

Figure 3. Map of main track-bearing surface with sauropodomorph trackways JYS1-10.

4 L. Xing et al.884

trackmaker. Tracings were also used to make overlay

images, as proposed by Baird (1957). In this method,

tracings of multiple tracks from the same trackway were

overlain to give a composite, idealised image. A replica of

a pes imprint UCM 178.18 was made using latex.

4. Description of the trackways

The trackways are conspicuous and generally deep and

continuous over much of the main surface (Figures 2 and 3).

The majority occur on a surface within the upper sandstone

unit, but one easily accessible trackway occurs on a surface

one metre above the main surface on the uppermost surface of

the upper sandstone bed (Figure 4). There are at least two

trackway morphotypes present. The first morphotype (A),

found on the main track-bearing surface, is characterised by an

oval, outwardly rotated pes with four pes claw traces sub-

parallel to the footprint axis, and a semicircular to pentadactyl

manus (Figures 5–9). The second morphotype (B) is a more

typical sauropod morphotype characterised by an oval to sub-

triangular, outwardly rotated pes, some with outwardly rotated

claw traces, and a semicircular, outwardly rotated manus

(Figures 10–11). A good example of this morphotype is the

trackway found on the upper surface, just above the main track

level. Some of these differences may be due to variation in

trackmaker size and quality of trackway preservation.

Figure 4. Stratigraphic section showing position of upper track-bearing sandstone unit and other track-bearing levels.

Historical Biology 5885

4.1. Systematic paleontology

Liujianpus ichnogen nov. Figures 5–9

Type ichnospecies: Liujianpus shunan (monospecific)

Horizon and locality: Daanzhai Member of Ziliujing

Formation (Lower Jurassic), Jiaoyuan near Gulin County,

Sichuan Province China (latitude N 248 020 32.7800 andlongitude E 1058 48053.0500).

Etymology: after LIU Jian, a famous Chinese

mountaineer and explorer, who helped with technical

climbing at this and other sites, and pus meaning ‘foot’.

Diagnosis. Trackway of large quadruped with

elongate, tetradactyl, outwardly rotated pes with length

of digit III . IV ¼ II . I and a sole area divided by atransverse crease. Manus pentadactyl to semi-circular and

strongly rotated outward. Differs from other sauropodo-

morph tracks by the presence of four nearly equidimen-

sional, elongate pes digit traces oriented subparallel to the

pes axis and a sub-circular manus with five discrete blunt

digit traces. Combines some features of otozoid tracks

with those of sauropod tracks such as Brontopodus.

Liujianpus shunan ichnosp nov

Holotype: Trackway JYS1 comprising 15 pes and

14 manus tracks including JYS1-LP7 and LM7 (replica

UCM 178.18) (Figures 3, 5–8, 9A) (Table 1).

Paratype: Trackway JYS3 with 16 pes and 15 manus

tracks (Figures 3, 5, 6–9B) (Table 2).

Horizon and locality: As for the ichnogenus.

Etymology: the specific name ‘shunan’ refers the

southern part of the Sichuan Province.

Diagnosis: As for the ichnogenus.

Figure 5. Photograph (A) and interpretative outline drawing (B) of trackways JYS1 (holotype) and JYS3 (paratype).

6 L. Xing et al.886

4.1.1. Description

The pes imprints are tetradactyl with their length (l) and

width (w) ranging from about 37–40 cm and 26–28 cm,

respectively (average l/w ratios 1.32–1.54). They are

outward-rotated, terminating anteriorly in four equidimen-

sional, slightly curved digit traces that are oriented sub

parallel to the pes axis (Figures 5–9). Pes sole area divided

by a transverse crease, situated almost midway along the

length of the tracks: i.e. about 40–45% of pes length from

the anterior margin of the footprint. On the medial border,

this crease terminates in a distinctive notch behind digit

I. Pes claw traces make up about 25–30% of pes length.

Figure 7. Photographs of pes tracks JYS1-LP7, LP8; JYS3-LP1.

Figure 6. Photographs and interpretative outline drawing of manus-pes tracks JYS1-LP7, LM7 (holotype), JYS3-RP2, RM2 and JYS3-RP3, RM3 (paratype).

Historical Biology 7887

Manus pentadactyl to semi-circular, wider than long,

with length (l) between 15 and 18 cm and width (w) about

25 cm, (l/w ratios 0.60–0.74), subcircular to pentadactyl in

outline.

Trackway narrow with pes inner trackway width

6–12 cm and pes outer trackway width ,75 cm. Innertrackway width greater for manus than for pes (,24–28 cm), but outer trackway width slightly less (63–70 cm).

Outward rotation of the pes averages 38–408, outwardrotation of the manus ,528. Pes pace angulation averagesbetween 112 and 1178, manus pace angulation averages105–1118. Pes step and stride averaging about 70–86 cmand 120–145 cm, respectively, manus step and stride

averaging about 75–77 cm and 118–123 cm, respectively.

4.1.2. Brontopodus-type

An isolated trackway (JYS11) (Figure 10) of a small

sauropodomorph from the upper surface of the upper

sandstone appears to show all the features of a typical

sauropod trackway such as Brontopodus-type (Table 3),

except that it is relatively narrow gauge. It is small, with a

mean pes length and width of 26.4 and 18.7 cm,

respectively (l/w 1.41) and corresponding mean manus

length and width of 10.4 and 18.3, respectively (l/w 0.57),

mean outward rotation for the pes is 348 and mean paceangulation is 1098. These values are all similar to thoserecorded for trackways JYS1 and JYS3 from the main

track-bearing surface (Tables 1 and 2). However, these are

generalised, non-diagnostic features. Trackway JYS11

shows almost no pes or manus digit traces except in a fewFigure 9. Composite manus pes outlines based on trackwaysJYS1 (A, holotype) and JYS3 (B, paratype).

Figure 8. The 3D height map (A), and interpretative outline drawing (B) of the JYS3-LP3 and RM3.

8 L. Xing et al.888

cases where the trace of a manus pollex (digit I) appears to

be present.

An easily accessible small pes JYSI-1 (Figure 11) with

a maximum length of 26 cm and width of 18 cm (l/w 1.44)

is exposed on the lower part of the main track surface in

the eastern sector of the outcrop. This track was replicated

and is preserved as UCM 178.19. The track shows strong

outward rotation of the traces of digits I–III, but the trace

of digit III is inconspicuous and there is no clear trace of

digits IV and V. The trackway is incomplete. As the JYS11

trackway, JYSI-1 can be assigned to the Brontopodus-type

track.

5. Discussion

The diagnostic features of Liujianpus are the four elongate

slightly curved, equidimensional pes claw traces aligned

sub-parallel to the pes axis and the pentadactyl blunt

manus. The relative lengths of the pes digit traces

III . IV ¼ II . I are also diagnostic and can be shown torecur in consecutive right and left footprints in multiple

trackways (Figures 5–9). Likewise, the distinctive

transverse crease, ending medially in a notch behind the

trace of digit I on the pes, is distinctive and seen in

multiple tracks. Such a combination of features has not

been reported from other sauropodomorph footprints,

except for those in Ishigaki (1988).

Thus, the closest match for the tracks from Jiaoyuan

are those illustrated by Ishigaki (1988, Figures 11 and 23)

from the reliably dated Lower Jurassic Assif n Sremt

(Sremt Assifn) site in Morocco. Although these tracks

show the notch behind digit 1 quite clearly, Ishigaki (1988)

did not describe these tracks in detail. Based on the

illustrations: (1) the Moroccan tracks represent a

quadruped; (2) they represent large animals (pes length

and width 75 and 61 cm, respectively) making them almost

twice the size of the Sichuan tracks; (3) the outward

rotation of the pes is similar as are the relative digit lengths

(III . IV ¼ II . 1) and (4) the medial notch behind pesdigit I seems to divide the track transversely as in the

Sichuan footprints. The most significant difference is that

the manus in the Moroccan trackways is less outwardly

rotated and relatively smaller and divided into two

impressions giving it a bilobed shape.

Otozoum from the Early Jurassic of North America

(Hitchcock 1847, 1848, 1858; Lull 1904, 1953) has several

similarities to Liujianpus, most notably in the relative

lengths of the pes digits (III . IV ¼ II . I). However,unlike Liujianpus, Otozoum has highly segmented pes

digits with as many as three digital pads on the distal

portions of the digits, which are clearly separated by the

interdigital hypices that extend posteriorly for as much as

40–50% of pes length. Also, the inner digit traces of

Liujianpus (digits I and II) are longer: i.e. more anteriorly

Table 1. Measurements for Liujianpus holotype trackway JYS1. Specimens JYS1-LP7 and LM7 are holotype.

Track ID Length Width Pes rot Pace angle Step Stride

JYS1-RP7 36.0 28.5 408 – – –JYS1-RM7 17.0 25.0 658 – – –JYS1-LP7 42.0 27.5 408 1108 90.0 143.0JYS1-LM7 24.0 25.5 458 1058 86.0 128.0JYS1-RP8 33.5 29.0 328 1248 82.0 146.0JYS1-RM8 14.5 24.5 618 1058 75.0 118.0JYS1-LP8 37.0 27.5 308 – 88.0 –JYS1-LM8 17.5 24.0 378 – 71.0 –Mean-P 37.1 28.1 388 1178 86.0 144.5Mean-M 18.3 24.6 528 1058 77.3 123.0

Table 2. Measurements for Liujianpus paratype trackway JYS3.

Track ID Length Width Pes rot Pace angle Step Stride

JYS3-LP1 40.5 26.0 428 – – –JYS3-LM1 16.5 24.0 628 – – –JYS3-RP2 42.5 27.0 408 1108 65.5 124.0JYS3-RM2 12.0 22.0 588 1108 75.5 120.0JYS3-LP2 38.0 27.0 458 1128 84.5 126.0JYS3-LM2 13.5 25.0 688 1128 76.0 115.0JYS3-RP3 40.0 26.5 308 1148 65.0 109.0JYS3-RM3 18.0 29.5 648 – 75.5 –JYS3-LP3 40.0 24.0 438 – 65.0 –Mean-P 40.2 26.1 408 1128 70.0 119.7Mean-M 15.0 25.1 638 1118 75.5 117.5

Historical Biology 9889

developed than in Otozoum, and the pes is far more

strongly outwardly rotated. Otozoum trackways indicate

bipedal progression except in rare cases including the

holotype (Lull 1953; Rainforth 2003; Lockley et al.

2006a). Moreover, the rarely preserved manus is small in

comparison with Liujianpus, more gracile and has well-

separated digit traces. Also, in Otozoum, it is positioned

laterally rather than anteriorly to the pes. Avanzini et al.

(2003) claimed that Lavinipes was very similar to

Otozoum, except that it evidently represented a fully

Figure 10. Photograph (A) and interpretative outline drawing (B) of small sauropod trackway JYS11 from the Jiaoyuan tracksite.

10 L. Xing et al.890

quadrupedal trackmaker. In this regard, Liujianpus is

closer to Lavinipes in representing a quadruped with a

manus larger than in Otozoum. However, the differences in

the pes digit size and outward rotation between Liujianpus

and Lavinipes are just as pronounced as those outlined

earlier between Liujianpus and Otozoum.

Liujianpus is similar to typical Brontopodus or

Brontopodus-like sauropod trackways in a number of

general, mostly non-diagnostic, features. Most notable are

the similarities in the oval, elephantine, outwardly rotated,

fleshy pes, with large anterior claw traces and the

semicircular outwardly rotated manus. Pace angulation

values are similar to those reported for typical sauropod

trackways (vid. Farlow et al. 1989). However, the digit

configuration and orientation of Liujianpus is quite

different. Notably, digit I is the shortest and digit III the

longest, with digit IV being well developed. This is in

contrast to the typical sauropod configuration where digit I

is the longest, with digits II–V being progressively shorter

(I . II . III . IV . V). In Liujianpus, digit I is lessconspicuous, and, contrary to Brontopodus, digits IVand V

lack claw traces. The different configuration III . IV ¼II . I of digit traces and their orientation subparallel to thefootprint long-axis strongly supports a distinct ichnogenus

Liujiangpus.

Although some of the Liujianpus manus tracks are

semicircular to crescent shaped without distinct digit

traces, unlike many sauropod tracks, a significant number

show distinct traces of all five manus digits (Figure 5).

Such configurations are rare in sauropod tracks and are

again more reminiscent of the basal sauropodomorph track

Otozoum. However, as noted earlier, Otozoum manus

tracks are rare and smaller, with more clearly differ-

entiated digits yhasn in Liujianpus.

Figure 11. Photograph (A) and outline drawing (B) of small sauropod pes track JYSI-1 from the Jiaoyuan tracksite.

Table 3. Measurements for sauropod trackway JYS11.

Track ID Length Width Pes rot Pace angle Step Stride

JYS11-RM1 10.5 18.5 – – – –JYS11-LP1 24.5 19.0 358 – – –JYS11-LM1 10.5 20.0 – – – –JYS11-RP2 27.0 18.5 358 1058 66.0 105.0JYS11-RM2 9.0 18.5 – – – –JYS11-lp2 29.5 19.0 448 1078 66.0 109.0JYS11-lm2 12.0 20.0 – – – –JYS11-rp3 25.0 19.0 178 1128 65.0 116.0JYS11-rm3 10.5 17.0 – – – –JYS11-lp3 25.5 17.5 308 1128 69.0 110.0JYS11-lm3 10.5 19.0 – – – –JYS11-rp4 27.0 19.0 43 1098 59.0 96.0JYS11-rm4 10.0 15.0 – – – –JYS11-lp4 26.0 19.0 408 – 55.0 –Mean-P 26.4 18.7 348 1098 63.3 107.2Mean-M 10.4 18.3 – – – –

Historical Biology 11891

Liujianpus shows a combination of features shared

with both Otozoum and Brontopodus. Therefore, it cannot

be assigned to either of them, and is considered here as a

new ichnogenus (Table 4). To date, the closest match for

overall trackway configuration and pes track morphology

comes from the unnamed Moroccan material reported by

Ishigaki (1988).

The discovery of a large sample of sauropodomorph

tracks combining morphological features of well-known

basal sauropodomorph and sauropod ichnogenera indi-

cates that the morphological diversity of sauropodomorph

tracks, and their relationship to the evolution of the

sauropodomorph pes, is still not fully understood.

Moreover, the occurrence of such intermediate mor-

phologies has implications for the evolution and

diversification of sauropodomorphs, especially in the

early Mesozoic.

Fortunately, sauropodomorph dinosaurs are well

known in southern China from the early Jurassic. For

example, the basal sauropodomorph Lufengosaurus

(Young 1941) is abundant in Yunnan Province, and its

pedal morphology (Figure 12) is consistent with the

Liujianpus morphotype. Lufengosaurus co-occurs with

other basal sauropodomorph such as Yunnanosaurus

huangi and various Early Jurassic sauropods such as

Chinshakiangosaurus and Jingshanosaurus. According to

Upchurch et al. (2007), Chinshakiangosaurus possesses an

unusual combination of basal sauropodomorph and

sauropod features. Thus, the Jiaoyuan footprints seem

also just as useful as skeletal remains in inferring

intermediate or mixed characteristics of early sauropodo-

morph morphology in the Early Jurassic of Asia.

Remarkably, some complete sauropodomorph skel-

etons were discovered in the Lower Jurassic Dongyuemiao

Member of the Ziliujing Formation in Gongxian County in

the southern Sichuan Basin, which were named Gongxia-

nosaurus shibeiensis (He et al. 1998). Gongxianosaurus is

approximately 14 metres in body length. He et al. (1998)

and Luo and Wang (2000) assigned the specimens to early

sauropods according to the ratio of fore limbs to hind limbs

was 70–75%. The Gongxian specimens were assigned to

basal sauropodomorph based on characteristics such as the

amphiplatyan–amphicoelous vertebrae and an ilium with

typical basal sauropodomorph morphology. The phalan-

geal formula is 2-3-4-5-1 (Dong Z.M. pers. comm.).

A recent analysis by Apaldetti et al. (2011) suggests that

Gongxianosaurus was more basal than Vulcanodon,

Tazoudasaurus and Isanosaurus, but more derived than

the early sauropods Antetonitrus, Lessemsaurus, Blikana-

Table 4. Comparison between Otozoum, Liujianpus and Brontopodus indicates which features are characteristic of prosauropods (p) orsauropods (s).

Ichnogenus Trackway patternGeneral pesshape

Pes digittraces

Pes digitlengths Pes–manus rotations Manus outline

Manus digittraces

Otozoum p p p p p p pLiujianpus s s p p s p/s pBrontopodus s s s s s s s

Note: The diagnostic morphological features of Liujianpus are equally divided between those characteristic of prosauropods and those typical ofprosauropods.

Figure 12. Well-preserved Lufengosaurus huenei pes skeletonZLJ 0023 from Lower Jurassic Lufeng Basin, Yunnan Provincefor comparison.

12 L. Xing et al.892

saurus, Camelotia and Melanorosaurus. In general, the

combined characteristics of Basal sauropodomorph and

Sauropoda are reflected in Gongxianosaurus, just as they

are in Liujianpus.

Most sauropod trackways in China are wide- (or

medium-) gauge and are therefore referred to the

ichnogenus Brontopodus (Lockley et al., 2002). Xing

et al. (2010, 2013, 2015) also assigned the Early

Cretaceous narrow-gauge, medium-sized/small-sized

sauropod trackways from the Nanguzhai tracksite, Jishan

tracksite and the Qianjiadian tracksite to Parabrontopodus

isp. However, the Jiaoyuan JYS11 trackway lacks a strong

outward rotation of the manus, which is a typical

characteristic in Chinese medium-sized/small-sized Para-

brontopodus tracks (Xing, Lockley, Bonnan et al in press).

Therefore, the characteristics of Jiaoyuan JYS11 trackway

are more closer to the Brontopodus-type tracks. Lastly, the

presence of a small Brontopodus-like trackway in close

association with the assemblage of much larger Liujianpus

indicates that small and large sauropodomorphs may have

co-existed.

6. Conclusion

1. The Jiaoyuan track site reveals the largest

sauropodomorph trackway assemblage currently

known from the Early Mesozoic (probably Lower

Jurassic) of China, and indeed from all of Asia.

2. The site consists of multiple sub parallel and

bimodally oriented trackways on a steep, near

vertical wall.

3. The trackways include at least two morphotypes:

a Brontopodus-type form of presumed ‘true’

sauropod affinity, and a new sauropodomorph

ichnotaxon, Liujianpus shunan ichnogen. nov.

ichnosp. nov. of probable basal sauropodomorph

affinity.

4. Liujianpus combines features of Brontopodus,

Otozoum and the Otozoum-like European sauropo-

domorph track Lavinipes, but is morphologically

distinct from all three in pes digit and manus track

morphology.

5. It is somewhat similar to an unnamed Early Jurassic

track from Morocco.

6. The combination of morphological features attrib-

uted to both basal sauropodomorph and sauropods

expands the range of sauropodomorph known

footprint morphologies.

7. The footprint morphotypes may be linked with foot

morphologies intermediate between basal sauropo-

domorph and sauropods, which is consistent with

the skeletal record of sauropodomorphs in the Early

Jurassic of Asia.

Acknowledgements

The authors thank Jian LIU and Bing Xiao (SichuanMountaineering Association, China) for their participation infield research. Detailed revisions by Gerard Gierliński, JeremyLockwood and Stu Pond greatly improved the final version andwe are appreciative of their efforts.

Disclosure statement

No potential conflict of interest was reported by the author(s).

Funding

This work was supported by the special projects grants ofZhaojue County, by the special projects grants of Gulin CountyPeople’s Government, Sichuan Province, by the special projectsgrants of Yanqing Geopark, Yanqing County, Beijing, and by the2013 support fund for graduate students’ science and technologyinnovation from China University of Geosciences (Beijing),China.

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Abstract1. Introduction1.1. Global distribution of basal sauropodomorph tracks1.2. Sauropodomorph trackways from Gulin County

2. Geographic and geologic setting3. Methods4. Description of the trackways4.1. Systematic paleontologyLiujianpus ichnogen nov. Figures 5-9 Liujianpus shunan ichnosp nov4.1.1. Description4.1.2. Brontopodus-type

5. Discussion6. ConclusionAcknowledgementsDisclosure statementFundingReferences